The photoreceptor and PAS/LOV protein VIVID (VVD) modulates blue-light signaling and influences light and temperature responses of the circadian clock in Neurospora crassa. One of the main actions of VVD on the circadian clock is to influence circadian clock phase by regulating levels of the transcripts encoded by the central clock gene frequency (frq). How this regulation is achieved is unknown. Here we show that VVD interacts with complexes central for circadian clock and blue-light signaling, namely the WHITE-COLLAR complex (WCC) and FREQUENCY-interacting RNA helicase (FRH), a component that complexes with FRQ to mediate negative feedback control in Neurospora. VVD interacts with FRH in the absence of WCC and FRQ but does not seem to control the exosome-mediated negative feedback loop. Instead, VVD acts to modulate the transcriptional activity of the WCC.ight, in addition to providing an energy source for many life forms on Earth, acts as a signal that may trigger development or serve as a repetitive cue that marks the passing of external time. External time cues are used by cellular timers such as circadian clocks to lock their periods to that of the external day. The process of period locking is called "entrainment" and ensures that cellular and behavioral activities happen at times of day when their adaptive value is highest (1-3). Blue light plays a central role in the entrainment of circadian clocks. Indeed blue-light photoreceptors and circadian clocks may have coevolved from a mechanism that originally served to detect (photoreceptor) and avoid (timer) harmful radiation (4-6). Our understanding of the molecular bases of circadian clocks and their responses to light has improved dramatically during the last decade or so, and the eukaryotic model organism Neurospora crassa has become one of the best-studied systems for understanding both processes (7-9).The key components of the Neurospora circadian clock are the products of the white collar (wc-1 and wc-2), frequency (frq), and frq-interacting helicase (frh) genes (4, 10, 11). The blue-light photoreceptor WC-1, and its interaction partner WC-2, form the transcriptionally and photoactive WHITE COLLAR complex (WCC) that activates frq expression (4, 12). FRQ protein, in turn, complexes with FRH to form an FRQ-FRH complex (FFC) that represses WCC activity (9, 11). Thus, photoreception and temporal organization of gene expression are linked via the WCC (4,(12)(13)(14). Hyperphosphorylated WCC is transcriptionally less active, and repression of WCC by FRQ occurs via FRQ-mediated phosphorylation of WCC by Casein kinase 1 and 2 (CK1 and 2) (14, 15).A second feedback loop that acts to repress WCC activity involves the product of the vivid (vvd) gene (16). Like WC-1, VVD is a PAS/LOV protein and blue-light photoreceptor; however, unlike WC-1, its presence is not essential for circadian rhythmicity in constant darkness (DD) (16)(17)(18)(19). Nevertheless, VVD has important roles within the Neurospora circadian system. Without VVD the organism is more sensitive to ...